A Resilience Framework
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CHAPTER THIRTEEN Disease Effects on Landscape and Regional Systems: A Resilience Framework F. Stuart Chapin III, Valerie T. Eviner, Lee M. Talbot, Bruce A. Wilcox, Dawn R. Magness, Carol A. Brewer, and Daniel S. Keebler Summary In this chapter we present and evaluate a conceptual framework in- tended to improve predictability of the role of disease in landscape pro- cesses of socioecological systems. On a local scale, disease tends to increase the interactions among patches on a landscape, particularly through changes in the frequency and severity of disturbances and increases in the vulnerability of ecosystems to fundamental changes in state in re- sponse to such disturbances. In addition, human activities that alter landscape structure or connectivity often increase the likelihood of dis- ease epidemics or landscape sensitivity to disease. In contrast, on global and regional scales, disease tends to reduce landscape connectivity be- cause the social response to these ecological changes is often to restrict trade and migration. Both the increase in connectivity on the local scale and the decrease in connectivity on regional to global scales can pro- foundly affect ecosystem structure and functioning. Resilience theory addresses the factors that make systems either resil- ient or vulnerable to radical changes in state by considering interactions between the ecological and social components of systems. This provides a framework for predicting the regional impacts of disease and reducing their social consequences. Resilience is facilitated by fostering cultural and ecological legacies that provide seeds for recovery; by maintaining biological, social, institutional, and economic diversity, which increases the options for alternative pathways of recovery; and by fostering learn- ing and innovation, which reduces the potential impacts of disease and increases the likelihood of favorable social change. Introduction Many of the key drivers of landscape and regional pro cesses are chang- -1— ing at unpre ce dented rates and in ways that are strongly infl uenced by 0— human activities (Foley et al. 2005; Steffen et al. 2004; Vitousek et al. +1— 34276_ch02.indd 284 8/17/07 3:59:36 PM EFFECTS ON LANDSCAPE AND REGIONAL SYSTEMS 285 1997). Diseases of plants, animals, and people often play a key role in these changes, both as an integral component of socioecological systems (Eviner and Likens, chapter 12, this volume) and as a trigger for changes in state. Yet disease is often viewed as a surprise that cannot be antici- pated and therefore is not incorporated into predictive pro cess- based frameworks. Earlier chapters in this book described the impacts of dis- ease on individual organisms, communities, and ecosystems. In this chapter we extend these effects to larger spatial scales—landscapes and regions. We fi rst assess the role of disease in landscape processes through a conceptual framework that relies on the integration of social and eco- logical systems. Human activities are an integral component of system dynamics on regional scales, as delineated in previous Cary Conferences (Groffman and Likens 1994; McDonnell and Pickett 1993), so we ex- plicitly consider the social and ecological pro cesses that characterize the reciprocal interactions of socioecological systems (Berkes et al. 2003; Machlis et al. 1997) (fi gure 13.1) and determine the resilience of these Figure 13.1. A socioecological system. The oval represents the socioecological system and the circles delineate its ecological and social subsystems. Several in- de pen dent factors, such as climate, history, and international markets, deter- mine the properties of the system; these properties are further modifi ed by interactive controls that determine the internal dynamics of the system. (Re- —-1 printed with permission from Whiteman et al. 2004.) — 0 —+1 34276_ch02.indd 285 8/17/07 3:59:36 PM 286 CHAPIN ET AL. systems. Resilience is the capacity of a system to absorb shocks such as disease outbreaks without changing its fundamental properties, such as its social norms, its typical economy, and the types of species it supports (e.g., grain crops or forests). Using this framework, we describe the ef- fects of disease on landscape pro cesses. We then use resilience theory to explore the role of disease and insect pests in causing changes in the state of socioecological systems, using bark beetles, rinderpest, malaria, and AIDS as examples. Conceptual Framework Pathogen and parasite interactions with their hosts are such an integral component of community dynamics that their chronic role in ecosys- tems is nearly invisible (Horwitz and Wilcox 2005; Lafferty, chapter 9, Eviner and Likens, chapter 12, this volume). As discussed in earlier chapters, diseases often modify competitive interactions, trophic dy- namics, disturbance probabilities, and succession. Disease epidemics emerge when the pathogen- host balance shifts to be seriously deleteri- ous to the host (Horwitz and Wilcox 2005), often creating unantici- pated surprises (Gunderson 2003) that act as disturbances to alter the structure, composition, and functioning of ecosystems and landscapes (Wilcox and Gubler 2005). Many disease epidemics are triggered by human- induced environmental or biotic changes that shift the host- pathogen balance (Patz 2002; Patz et al. 2000). Here we examine dis- ease from this perspective, considering its role in ecosystems and how ecosystem resilience infl uences and is infl uenced by disease events. Diseases with the greatest immediate ecosystem impacts are generally those that selectively remove or suppress the dominant and keystone plant species (fi gure 13.2), which, by defi nition, are the plant species with the greatest effects on ecosystem pro cesses (Chapin et al. 2002). Diseases can have equally dramatic impacts by selectively removing ani- mals that control the abundance of dominant and keystone plants through trophic cascades (Polis 1999). In many aquatic ecosystems, bac- teriophages (viruses that attack decomposer organisms) have a profound immediate direct effect on nutrient cycling by stimulating bacterial turn- over (Middelboe, chapter 11, this volume). Over longer time scales, dis- eases infl uence ecosystem dynamics through their effects on organisms that infl uence disturbance regimes or species interactions (Eviner and Chapin 2003; Eviner and Likens, chapter 12, this volume). Disease has equally profound effects on the human component of so- -1— cioecological systems (fi gure 13.3). Disease has its greatest effects on so- 0— cial systems when it alters human population densities and activities, +1— 34276_ch02.indd 286 8/17/07 3:59:38 PM cesses. Major ecological pathways by which disease outbreaks affect landscape pro Figure 13.2. —-1 — 0 —+1 34276_ch02.indd 287 8/17/07 3:59:38 PM cesses. The Major social pathways by which disease outbreaks affect landscape pro Figure 13.3. effects of disease on social responses are largely mediated by learning and education. and learning by mediated largely are responses social on disease of effects -1— 0— +1— 34276_ch02.indd 288 8/17/07 3:59:40 PM EFFECTS ON LANDSCAPE AND REGIONAL SYSTEMS 289 either by directly infecting humans (e.g., bubonic plague, Yersinia pes- tis, malaria, Plasmodium, tuberculosis, Mycobacterium tuberculosis, sleeping sickness, Trypanosoma brucei rhodesiense and T. b. gambiense) or by disrupting the food supply (e.g., the Irish famine precipitated by potato late blight, Phytophthora infestans, which led to the death of more than a million people and the migration of a million more) (Mc- Neil 1989). Pathogens exert many other effects, including those that al- ter ecosystem and landscape pro cesses (see fi gure 13.2) in ways that affect the delivery of ecosystem ser vices to society (Millennium Ecosys- tem Assessment 2005). Human diseases often have differential effects on partic u lar segments of society (Turshen 1977). Those diseases that directly affect strong eco- logical agents, such as farmers, foresters, or pastoralists, have dispro- portionately large ecological consequences. Malaria and schistosomiasis, for example, have their greatest impacts on rural populations and there- fore on agricultural land use (Sachs and Malaney 2002). In addition, diseases that are perceived as posing a general health risk (particularly a risk to privileged segments of society) are more likely to infl uence policy than are diseases that are largely restricted to disenfranchised popula- tions (Turshen 1977). These policies in turn often infl uence public health decisions, land and water management, travel, trade, and other human activities. AIDS, for example, initially received only modest research funding in the United States because it was viewed primarily as a disease of homosexuals. It received more attention by policy makers after it be- came recognized as a health risk for broad segments of society. In con- trast, severe acute respiratory syndrome (SARS), which was immediately perceived as a health risk for the general public, particularly global trav- elers, received more immediate control efforts. Plant and animal diseases also have important consequences that are mediated by socioecological feedbacks (fi gure 13.3). Eco nom ical ly im- portant diseases (e.g., foot- and- mouth disease, Aphthovirus, soybean rust, Phakopsora, white pine blister rust, Cronartium ribicola) decrease the productivity of hosts that support the human food and fi ber supply and the stability of economic and social systems. These diseases often trigger management actions that remove diseased organisms or alter landscape linkages in an attempt to isolate areas with the disease. An outbreak of foot- and- mouth disease in England, for example, reduced beef exports and severely restricted regional travel in 76% of the coun- try. The restriction on the movement of cattle led to overgrazing of pas- tures (Fraser of Allander Institute [FAI] 2005). On the fl ip side, the establishment of Uruguay as a foot- and- mouth disease- free country led to a doubling of its beef exports and a corresponding expansion of pas- —-1 turelands (Food and Agriculture Or ga ni za tion [FAO] 2001). In this — 0 —+1 34276_ch02.indd 289 8/17/07 3:59:41 PM 290 CHAPIN ET AL.